Shane Et Al

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MARINE MAMMAL SCIENCE, 2(1):34-63 (January 1986)

0 1986 by the Society for Marine Mammalogy
ECOLOGY, BEHAVIOR AND SOCIAL

ORGANIZATION OF THE BOTTLENOSE
DOLPHIN: A REVIEW
SUSAN H. SHANE
RANDALL S. WELLS
Center for Marine Studies, Long Marine Laboratory, University of California,
Santa Cruz, California 95064
BERND WüRSIG
Moss Landing Marine Laboratories, P.O. Box 450, Moss Landing, California 95039
The authors review the literature on bottlenose dolphin ecology, behavior

and social organization, focusing on data collected on free-ranging animals. Most
bottlenose dolphins studied to date have had definable home ranges, and be-
havioral, morphological and biochemical information indicates discrete stocks
in some areas. Bottlenose dolphins appear to form relatively permanent social
groups based on sex and age. Mother-calf bonds are long-lasting. Movement
patterns are extremely variable from location to location but are relatively pre-
dictable at any given site. Food resources are one of the most important factors
affecting movements. Bottlenose dolphin behavior is very flexible, and these
dolphins are generally active day and night. Feeding peaks in the morning and
afternoon have been observed at several sites. Social behavior is an important
component of daily activities. Sharks are the most significant predator on bot-
tlenose dolphins in most areas, but captive and wild studies show that dolphins
and sharks frequently live in harmony as well. Human activities may be helpful,
harmful or neutral to bottlenose dolphins, but interactions with humans are
frequent for these coastal cetaceans.
Key words: bottlenose dolphin, Tursiops, ecology, behavior, social organization.
During the past 10-I 5 yr our knowledge of the behavior, social organization
and ecology of the bottlenose dolphin, particularly in the wild, has expanded
significantly. The primarily coastal nature of Tursiops truncatus and the wide-
spread use of this dolphin in oceanaria make it one of the best known of all
marine mammals. Advancing technologies suggest that we will be able to wrest
increasingly detailed information from free-ranging bottlenose dolphins. There-
fore, we have taken this opportunity to take stock of our present knowledge in
these areas.
34
SHANE ET AL.: BOTTLENOSE DOLPHINS: A REVIEW 35
HOME RANGE AND STOCK DISCRETENESS
The concensus is that there is one species of Tursiops worldwide, separated

into geographical races (Mitchell 1975, Rice 1977, Leatherwood and Reeves
1982). Distinctive inshore and offshore forms are typically present in most parts
of the bottlenose dolphin’s range (Caldwell and Caldwell 1972a, Walker 1975,
198 1, Leatherwood and Reeves 1982). It is clear that at least some coastal
bottlenose dolphins maintain home ranges. Researchers have identified both
individual and herd home ranges as well as apparently permanent and seasonal
home ranges.
Home Range
A home range is an area regularly used by an individual or group in the

course of performing normal daily activities (Burt 1943, Jewel1 1966). The
first indication that T. truncatus had a home range was provided by Caldwell
( 195 5), who defined a minimum home range for a recognizable individual
dolphin in Florida. Caldwell and Golley (1965) estimated a minimum home
range of “95 shoreline miles” for an albino bottlenose dolphin in Georgia and
South Carolina (previously described as a local animal by Essapian [1962)).
Caldwell and Caldwell (1972a) proposed that the species may have seasonal
home ranges linked by a traveling range.
The most detailed study of bottlenose dolphin home ranges has been con-
ducted on the west coast of Florida (Irvine and Wells 1972, Wells 1978, Wells
et al. 1980, 1981a, Irvine et al. 1981). The population of about 105 dolphins
appeared to maintain a home range of about 85 km*. Female-calf pairs and
subadult males had home ranges averaging approximately 40 km2, whereas
other adult females, subadult females and adult males had smaller ranges (15-
20 km2>. The dolphins used certain parts of the study area more during certain
seasons. For instance, they were seen more often in the passes and the Gulf of
Mexico during winter (Wells et al. 1980). In 1975-1976 Irvine et al. (1981)
recaptured 11 of 12 animals first taken in 1970-197 1 in the same area, and
Wells et al. ( 19836) reported that several individuals have been seen in this
area for more than 13 yr; this indicates that the home range is permanent.
Shane (1980) (also see Shane 1977, Shane and Schmidly 1978) found that
individual dolphins in her study area in Texas concentrated their activities in
certain areas, and she defined three major home ranges shared by several indi-
viduals. These ranges were used seasonally by some dolphins and year-round
by others. She believed the ranges of most dolphins extended outside of her
study area. Wiirsig and Wiirsig (1979) identified an apparent northern limit
for the Argentine population of T. truncatus they studied, but were unable to
determine how far the dolphins ranged beyond the 50 km* area they monitored.
However, they once sighted several known individuals 300 km away from their
study area. Gruber (1981) identified home ranges for individuals in her Texas
study area and reported on three nearly separate herd home ranges that coincided
partially with her study area. Hansen (1983) defined the normal range of a
population of bottlenose dolphins in southern California. Hussenot (1980)
36 MARINE MAMMAL SCIENCE, VOL. 2, NO. 1, 1986
reported that bottlenose dolphins found in the Molène archipelago of Brittany

had a larger range in winter than in summer. Saayman et al. (1973) found
that dolphins in Plettenberg Bay, South Africa, had a home range that extended
over at least 46 km.
Home range size may be a function of dolphin density, and variable densities
have been reported by different observers (Table 1). Some of this variability
may be due to different methods of observation and of calculating density, but
it may also indicate variable habitat quality.
Leatherwood (197 5) hypothesized that the wide range of feeding behaviors
exhibited by T. truncatus in different places represents a “plasticity” necessary
for animals with limited ranges and faced with changing food resources. Al-
though many bottlenose dolphins clearly concentrate their activities within home
ranges, how limiting these ranges are is unclear. Many studies show dolphins
changing their ranges seasonally: Mead (1975) described a seasonal migration
of bottlenose dolphins past Cape Hatteras, North Carolina. Wiirsig and Wiirsig
(1977) and Wiirsig (1978) found bottlenose dolphins capable of making a
600-km roundtrip away from what was thought to be their primary home
range.
Wells et al. (1980) provided information on the apparent exclusivity of
home ranges. While no defense of boundaries implying territoriality was ob-
served, dolphins in Sarasota seemed to recognize range limits and consistently
turned back at approximately the same location, thus defining the boundaries
of their range. T. truncatus in Brittany usually followed the same routes re-
peatedly (Hussenot 1980). Shane and Schmidly (1978), Shane (1980) and
Gruber (1981) noted an apparent boundary that was rarely crossed between .
inshore waters and the Gulf of Mexico at passes in Texas. Irvine et al. (198 1)
did not find the same limitation at passes on the Gulf coast of Florida although
the Sarasota dolphins were never seen more than 1 km offshore. Dolphins
presumably use underwater topography to recognize different locations within
their ranges (Würsig and Wiirsig 1979). Hansen (1983) implicated water
temperature as a possible factor limiting distribution in southern California.
Stock Discreteness
It is only within the past lo-15 yr that information contributing to the

identification of population stocks has become available. The degree of mixing
between populations can only be determined after individual population units
have been identified through consideration of (ideally) behavior, morphology
and biochemical genetic factors. Although complete data on these three aspects
are not available for any single population of bottlenose dolphins, a number of
recent studies have emphasized one or more of these aspects for different pop-
ulations.
The behavioral factors of primary importance in identifying populations of
bottlenose dolphins include movement and association patterns of individuals.
All of the studies of Tursiops movement patterns to date have dealt with coastal
populations; no data are available on the movements of bottlenose dolphins
Table 1. Bottlenose dolphin density reported in various studies.
Density
Location (dolphins/km2) Season Survey type Reference
Florida Gulf Coast 0.06 multiple aerial Leatherwood and Reeves 1982
Charlotte Harbor to Crystal River, FL 0.070 multiple aerial Odell and Reynolds 1980
Charlotte Harbor, FL 0.170 Oct. 1980 aerial Thompson 198 1
Charlotte Harbor, FL 0.206 Apr. 1981 aerial Thompson 198 1
North Gulf of Mexico 0.23-0.44 multiple aerial Leatherwood et al. 1978
Texas Gulf Coast 0.314 Sept. 1979 aerial Leatherwood and Reeves 1983
Charlotte Harbor, FL 0.469 Jan. 1981 aerial Thompson 198 1
Sarasota, FL (southern half of
study area) 0.6 multiple boat Wells 1978
Indian River, FL 0.68 Aug. 1977 aerial Leatherwood 1979
Texas Gulf Coast 0.75 Mar. 1978 aerial Barham et al. 1980
Pass Cavallo, TX 0.93 multiple boat Gruber 198 1
Corpus Christi Bay, TX 1.025 Sept. 1979 aerial Leatherwood and Reeves 1983
Indian and Banana Rivers, FL 1.22 Jan. 1980 aerial Leatherwood and Reeves 1982
Sarasota, FL 1.3 multiple boat Irvine et al. 1981
Aransas Pass, TX 1.4 Oct. 1976 boat Shane 1980
Sarasota, FL (northern half of
study area) 1.8 mutiple boat Wells 1978
Southern California 2.23-3.10 multiple boat Hansen 1983
Aransas Pass, TX 4.8 Jan. 1977 boat Shane 1980
more than a few kilometers from shore. In general, bottlenose dolphins in some
areas inhabit ranges that are common to entire groups and exclusive of other
groups (these may be considered population ranges); in other areas there may
be seasonal overlap in the ranges of individuals, and in still other areas dolphins
may make extensive migrations.
In many cases, consideration of individual ranges may be insufficient to define
populations behaviorally. In these cases, association patterns between individuals
can be helpful in resolving population memberships. For example, adjacent but
apparently separate populations have been noted in Texas and Florida. In Texas,
there was a lack of association between inshore and offshore dolphins off Aransas
Pass (Shane 1980) and Port O’Connor (Gruber 1981). In Florida a similar
lack of association was found between dolphins in the Gulf of Mexico or Tampa
Bay and those in inshore waters near Sarasota (Wells et al. 1980). Ten years
of observations of dolphins in the Sarasota area resulted in the behavioral
definition of a relatively discrete, resident population: 105 dolphins that regu-
larly moved through the same waters and interacted markedly more with each
other than with dolphins in adjacent areas (Wells et al. 1981a). Within the
85-km* population range of the Sarasota dolphins, 94 percent of the adult
females tagged during 1975-1976 were still present in 1982 while only one-
third of the males were seen.
This behavioral definition should not, however, be considered absolute for
the Sarasota population, nor should it be considered necessarily transferable to
other areas. On very few occasions, individuals or small groups from the Sarasota
population were observed with adjacent groups. In addition, four dolphins that
were not seen in the area for periods of several months or more were later
resighted in the area. Similarly, Wells et al. (1983a) reported a 1,500-km
roundtrip for identifiable dolphins first observed off southern California, and
Würsig (1978) described a 6OO-km roundtrip for six identifiable adult dolphins
over a 15-mo period off Argentina. Shane (1980) reported a sighting by Gruber
of an identifiable dolphin 100 km from its presumed home range. Asper and
Odell (1980) noted especially long movements by several of the dolphins they
tagged on the east coast of Florida. Lockyer (1978) described the movements
of an adult bottlenose dolphin off Great Britain as covering more than 500 km
in 18 mo. These exceptional movements, considered in light of the variable
dolphin densities in some areas, suggest that T. truncatus stocks are not abso-
lutely discrete; rather, the opportunity for mixing or genetic exchange between
populations exists unless there are as-yet-unknown social barriers between pop-
ulations that would preclude interactions. If mixing occurs between identifiable
populations, what is its extent. 2 This question cannot be approached solely
through consideration of behavioral studies; these studies need to be augmented
with more precise measures of the reproductive contribution of various members
or non-members to the population.
Studies of morphology and biochemical genetics of members of dolphin
populations, although more difficult to conduct because of the necessity of
hands-on sampling, allow the determination of population differences through
statistical comparison of a number of factors. Walker (1981) differentiated
SHANE ETAL.: BOTTLENOSE DOLPHINS: A REVIEW 39
between populations of bottlenose dolphins in the Pacific Ocean on the basis

of tooth size, skull length, size at sexual maturity, parasite loads and stomach
contents. Ross (1977) used similar criteria for animals off South Africa and
identified two forms. Another promising technique is the examination of blood
samples. Duffield (1980, 1981, 1982) has determined some degree of local
population differentiation by use of electrophoresis of blood proteins of dolphins
from the east coast of Florida. She also found a discernible pattern of interchange
between populations.
SOCIAL ORGANIZATION
The patterns of social organization of bottlenose dolphins appear to be quite

complex, based on detailed observations of captive dolphins supplemented by
information from a few studies of free-ranging animals. A first step in under-
standing social structure is to look at the size of dolphin groups.
Group Size
Group size is highly variable for bottlenose dolphins, due, in part, to differing
perceptions of the definition of a dolphin “group” (e.g., pod, herd, school,
subgroup, sighting) and different criteria used to determine membership in such
a unit. Group size ranges from one to more than 100, but bottlenose dolphins
are most commonly found in relatively small groups of 2-15 animals. One
notable exception is reported for bottlenose dolphins in South Africa where
group size ranged from 3 to 1,000 and averaged 140 per group (Saayman and
Tayler 1973). Wells et al. (1980, Table 6.2, pp. 276-277) and Leatherwood
and Reeves (1982, Table 18.2, p. 3 79) summarized information on group
sizes of bottlenose dolphins. Since these reviews, Leatherwood and Reeves (1983)
found mean group sizes of 6.10 and 5.2 3 dolphins in Corpus Christi Bay and
the Gulf of Mexico off southern Texas, respectively, and Hansen (1983) re-
ported a mean group size of 18 for bottlenose dolphins in coastal southern
California.
Wells et al. (1980) defined “primary groups” (= pods, Shane and Schmidly
1978, Gruber 1981) as the smallest units of dolphins that associated closely
and engaged in similar activities; these units were often intact for days or weeks
at a time. “Secondary groups” (= herds, Shane and Schmidly 1978, Gruber
1981) were temporary (minutes to hours) aggregations of primary groups (Ir-
vine et al. 198 1). Lear and Bryden (1980) described a similar grouping system
for bottlenose dolphins in Australia.
Habitat structure and activity patterns apparently are the prime factors in-
fluencing bottlenose dolphin group size. In general, group size tends to increase
with increased water depth or openness of the habitat. Wells (1978) reported
significantly larger groups in deep-water passes and the offshore Gulf of Mexico
than in the shallower inshore waters near Sarasota, Florida. Shane (1977) and
Gruber (1981) noted that groups were, on the average, larger when in open
waters than when in the constricted regions of channels or passes. Odell (1976)
found a similar pattern when he compared group size in the open waters of
Biscayne Bay to the complex aquatic habitat adjoining the Florida Everglades.
Leatherwood and Platter (1975) also reported larger groups in the open sound
of the northern Gulf of Mexico than in the shallow marshlands. Within the
marshlands, the largest groups were in deep channels connecting shallower
feeding areas (this relationship was not found in later work in the same region
reported by Leatherwood et al. 1978). Wiirsig (1978) noted that group size
averaged 14 near shore and 20 farther from shore. As discussed by Norris and
Dohl (1980a), Wells et al. (1980) and Wiirsig (1978), the reasons for these
variations in group size with physiography may be related to foraging techniques
and protection from predation. Shallow, inshore waters often provide relatively
predictable, evenly distributed food resources associated with reefs or seagrass
flats. In more open waters, schooling fish become the predominant resource
available to the dolphins. Larger groups of dolphins integrating their sensory
capabilities increase the probability of locating these patchy but rich food sources
and provide the numbers of individuals necessary to cooperatively locate and
herd prey. In the same way, larger groups in open waters serve some of the
same functions as the inshore physical habitat in terms of protection from
predation.
Variation in group size with time of day, reported by a number of researchers
(Shane 1977, Wells 1978, Wells et al. 198lS), is probably related to the
activity cycles of the animals. There are tendencies for groups of particular sizes
to be engaged in particular activities although these trends are not necessarily
consistent from location to location. Shane (1977) reported that groups engaged
in traveling, feeding and resting were approximately the same size and were
smaller than socializing groups. Lear and Bryden (1980) found idling (i.e.,
resting) groups to be smaller than swimming (i.e., traveling) groups. Wells,
Scott and Irvine (unpubl.) noted that traveling and socializing groups were
often smaller than feeding or resting groups.
Seasonal changes in group size have been reported, but they are inconsistent
from site to site. Odell and Reynolds (1980) noted an increase in group size
during the winter off Florida’s west coast, and Wiirsig (1978) observed the
same trend in Argentina. In contrast, Shane (1977) noted a slight decrease in
group size for bottlenose dolphins in Texas during winter, and Irvine et al.
(1981) found no significant seasonal variation in group size in bottlenose dol-
phins near Sarasota, Florida.
Social Structure: Captive Animals
Since 1940 studies of social behavior at a number of oceanaria have produced

relatively consistent results (e.g., McBride 1940, McBride and Hebb 1948,
McBride and Kritzler 19 5 1, Essapian 19 5 3, 1963, Tavolga and Essapian 19 5 7,
Tavolga 1966). The first intensively studied bottlenose dolphin colony, at what
is now Marineland of Florida, usually consisted of at least one mature male,
five or six mature females and their offspring and several subadults. This group
was structured into a hierarchy, with the largest adult male dominant over all
other tank-mates. A less rigid dominance hierarchy existed between the females,
with the largest females dominant over smaller animals (McBride and Hebb
1948, Tavolga 1966). Subsequent studies of colonies in Florida and elsewhere
suggest that dominance hierarchies may be the “typical” social system for
captive bottlenose dolphins although the hierarchy may not always be rigidly
maintained (Brown and Norris 1956, Caldwell et al. 1965, Tavolga 1966,
Caldwell and Caldwell 1967, 1972a, 6, Norris 1967, Saayman et al. 1973).
Dominance has been displayed by captive dolphins in the forms of jaw claps,
biting, ramming or tailslaps against subordinates (McBride and Hebb 1948,
Essapian 1953, Lawrence and Schevill 1954, Tavolga 1966, Norris 1967,
Caldwell et al. 1968, Caldwell and Caldwell 19726). During most of the year,
adult males either swam alone or for only brief periods with females; however,
longer associations, on the order of days or weeks, were maintained during a
courtship period, which was apparently terminated by the male (Essapian 19 5 3,
Tavolga and Essapian 1957, Tavolga 1966, Caldwell and Caldwell 1972b,
1977). Adult males captured from the same groups and maintained together
have been observed to maintain priority of access to food and females on the
basis of size of the male, but with little aggression (McBride 1940). Much of
the time, however, captive colonies have contained adult males from different
capture localities; in these cases the males have fought viciously during the
breeding season, to the extent that most oceanaria now generally maintain a
single adult male per tank (McBride and Kritzler 195 1, Wood 1977). This
suggests that dominance relationships may be long-established within dolphin
groups, with little need for frequent contests.
A variety of interactions between captive adult males and females with calves
have been reported. It is generally agreed that females without calves are the
preferred partners for adult males (Tavolga and Essapian 195 7, Caldwell and
Caldwell 1972a, S). There have been a number of reports of aggression, often
violent, by adult males toward calves, with the result being either reciprocated
aggression by the mother or prolonged avoidance of the adult males (McBride
and Hebb 1948, Essapian 1953, 1963, Tavolga and Essapian 1957, Caldwell
and Caldwell 1972a, d).
The dominance of adult males over subadult males has been frequently
reported as being expressed by aggression, especially when subadults were at-
tempting to copulate with females or when the younger animals were recently
added to an existing colony (McBride 1940, Tavolga 1966, Norris 1967,
Caldwell and Caldwell 1972a). The usual response by the subadults to this
aggression involved avoiding the adult males, often forming subadult male
groups.
Social Structure: Free-Ranging Animals
While the details of social interactions are more easily obtained under captive
conditions, the possible effects of this unnatural environment must be considered
when attempting to generalize the patterns of captives to free-ranging animals.
Field studies have shown that bottlenose dolphin group composition is much
more dynamic than previously believed (Wiirsig and Wiirsig 1977, 1979,
Wells 1978, Wiirsig 1978, Shane 1980, Wells et al. 1980, Irvine et al. 1981).
These researchers found that group composition changed frequently although
certain associations appeared to be more persistent or repeated more often than
others. Wells et al. (1980) reported that the associations of bottlenose dolphins
within a marked population on the west coast of Florida were based on the age
and sex of the individuals. In general, adult males formed small bands that
moved from female group to female group in one portion of the range of the
population, while subadult males swam in somewhat larger bachelor groups in
another portion of the range, occasionally moving with the female groups when
they passed through the area inhabited by the younger males. Adult and sub-
adult males were rarely seen together although their ranges overlapped slightly.
Continued observations of these marked animals 13 yr after initiation of the
marking studies have shown that these association patterns are recurrent and
long-lasting (Wells et al. 19836). Females with calves moved regularly through
apparent nursery areas (Irvine et al. 1981). Close associations between females
and calves were maintained for 3-6 yr, but one male was still with his mother
after at least 71/2--S yr (Wells et al. 198 lc, 19836). In western Australia bonds
between recognizable mothers and calves remained strong for 3-6 yr (Gawain
1984). This dependency period is longer than expected based on observations
of captive calves nursing for 18 mo or less (McBride and Kritzler 195 1). One
calf in a group of dolphins that frequent a beach in western Australia was
believed to nurse occasionally at 2ti yr of age (Gawain 1984). In Florida
associations between females tended to be based on the ages of their calves or
on the absence of calves, and female groups often lacked adult males (Wells et
al. 1981c, 19836). These observations suggest that male dominance is not the
overall controlling factor in the ordering of free-ranging social systems that it is
in captive situations.
While dominance may be expressed in a hierarchy in captitivy, it may be
expressed by positioning of individuals or subgroups and sexual segregation in
the wild (Norris and Dohl 1980a). Leatherwood (1977) noted that in the
northern Gulf of Mexico mothers and small calves were in the center of groups,
possibly as a protective mechanism against predation. The segregation of sub-
adult males as reported from the west coast of Florida has also been reported
from Cape Hatteras, North Carolina (True 1890, Townsend 1914). Conspecific
toothrakes have been observed more frequently on young males than other
classes in several cases in the Gulf of Mexico (Gunter 1942, Wells et. al.
1980). In the northern Gulf of Mexico, subgroups of young animals of un-
known sex were often observed near the periphery of large groups of bottlenose
dolphins (Leatherwood 19 77).
While the nature of the social organization of bottlenose dolphins remains

incompletely known, several generalizations can be made. At least in some
portions of the species’ range, populations form relatively permanent social units
that are closely tied to definable home ranges. Within these populations, social
associations and individual movements are based on the age and sex of the
individuals. In addition, the long-term associations between mothers and young,
and between other individuals, suggest that learning within the context of the
social unit is of major importance.
SEASONALAND DAILY MOVEMENTS
Seasonal Movements
While bottlenose dolphins may at times migrate in the higher-latitude ends

of their range (True 1890, Townsend 1914 and Mead 1975 provide descrip-
tions of movement past Cape Hatteras; see also Verwey 1975, Lockyer 1978
and Hussenot 1980 for seasonal movements in northern Europe), such move-
ments are often obscured by the fact that some animals remain in the same
areas year-round. This is particularly evident for populations at lower latitudes,
where “migrations ” as such cannot be discerned, but where there are definite
differences in numbers in different seasons. Thus, although Gunter (1942)
stated that there is no evidence for seasonal migration or seasonal change in
abundance of bottlenose dolphins in Texas waters, more detailed analyses showed
that there were twice as many dolphins present near Aransas Pass, Texas, during
the winter as during the summer (Shane and Schmidly 1978, Shane 1980).
Gruber (1981) found a similar trend for the Pass Cavallo area of Matagorda
Bay, Texas. Gruber found seasonal dolphin concentrations in association with
shrimp-fishing activities, but whether the movements of fish that were associated
with shrimp or the shrimpers’ operations were primarily responsible was not
determined. Caldwell and Caldwell (1972a) reported a limited seasonal move-
ment along the Atlantic coast of Florida, with some dolphins moving southward
in winter and northward in summer. However, these movements are probably
no greater than about 150 km (D. K. Caldwell, pers. comm., 1973). Irvine
and Wells (1972) reported no evidence for long-distance seasonal migrations
of dolphins in the Sarasota, Florida, area, and a subsequent analysis revealed
that in winter dolphins spent most of their time in passes and along the Gulf
coastline, while in summer they were found inshore of barrier islands (Irvine et
al. 1981). Furthermore, females and calves tended to aggregate over shallow
protected areas in summer, and dolphins tended to feed on mullet over shallow
flats from spring to fall. As mullet moved from shallow inshore waters to the
passes and into the Gulf of Mexico, dolphins also moved offshore and into the
passes, presumably adjusting to a change in this food supply (Wells et al.
1980, Irvine et al. 1981). Asper and Odell (1980) cited slight evidence that
dolphins may move further south from mideast Florida during winter, and
Odell (197 5, 1976) stated that a winter increase of animals in the Everglades
National Park may take place. However, Moore (1953) noted a lack of seasonal
migration in the Everglades. Lear and Bryden (1980) reported that numbers
of dolphins appear greater in southern Queensland, Australia, waters in winter
than in summer. Wiirsig and Wiirsig (1979) found bottlenose dolphins in
Argentina to be abundant except during the hottest months of the year. All of
44 MARINE MAMMAL SCIENCE. VOL. 2. NO. 1. 1986
these reports deal with bottlenose dolphins that are found close to land; little
is known about movements of the offshore populations.
In summary, some coastal bottlenose dolphins in higher latitudes show a
clear tendency towards seasonal migration, while those in warmer waters show
localized seasonal movements that probably have much to do with movements
of food and need for safety in reproduction. In at least one case, off western
Florida, movements of mullet are rather clearly correlated with bottlenose dol-
phin movements. The fact that bottlenose dolphins take advantage of many
different food items may explain why their movement patterns are not as clear-
cut as those of some other cetaceans.
Short-Term Movements
Diurnal and other short-term movement patterns appear to be influenced by

a variety of factors. Bottlenose dolphin movements have been described as
highly variable and flexible off Europe (Verwey 197 5, Duguy and Hussenot
1980). Lear and Bryden ( 1980) believed that bottlenose dolphins in eastern
Australia sought shelter near shore to avoid rough water offshore during storms.
They suggested that there was tidally related movement but not a simple
relationship with flood and ebb. In Argentina bottlenose dolphins moved into
deeper water during midday and behaved and fed in different ways in different
depths (Würsig and Würsig 1979). They also moved differently depending on
flood and ebb tides; such movement tended to keep the animals in shallower
water as the tide receded, until they were in water so shallow as to cause them
to abruptly move into deeper water. Dolphins off Argentina also moved back
and forth parallel to shore using underwater obstructions as cues to turn, thus
keeping them in a confined area of about 0.5 km distance along shore for
several hours before tidal changes or other factors caused them to abandon this
zig-zag pattern. Bottlenose dolphins off South Africa entered Plettenberg Bay
in the morning and afternoon, primarily to feed. It is likely that these animals
were following the diurnal cycles of several species of food fish, but the authors
did not detail the movements of prey (Saayman et al. 1973). Hoese (197 1)
reported dolphins entering salt marshes to chase fish onto mudbanks at low
tide. Caldwell and Caldwell (1972~) reported local movements of dolphins as
paralleling the northeastern coast of Florida, moving roughly southeasterly in
the morning and northwesterly in the afternoon. They hypothesized (as did
Pilleri and Knuckey 1968 for common dolphins in the Mediterranean Sea) that
this diurnal pattern may be sun-related.
Tidal flow often affects short-term movements. Bottlenose dolphins near
Sarasota, Florida, moved onto shallow seagrass flats with the incoming tide.
They fragmented into small groups during that time (Irvine and Wells 1972),
and much of the feeding was probably on mullet (Wells et al. 1980, Irvine et
al. 1981). Caldwell and Caldwell (1972a) also reported movement with tidal
flow in inland waters near St. Augustine, Florida. Shane (1980) described
movements in Aransas Pass, Texas, in which dolphins stationed themselves
against the tide, especially during resting. They showed a rough temporal pat-
SHANE ET AL.: BOTI'LENOSE DOLPHINS: A REVIEW 45
tern as well: in early morning, dolphins tended to move towards the north in
one part of the study area, during midday they moved in all directions, and
later they returned towards the south. Where the tide was strong, movement
against the tide was evident, but where the tide was weak, the temporal pattern
prevailed. Similarly, dolphins usually moved against the tidal flow near Pass
Cavallo, Texas (Gruber 1981).
Bottlenose dolphins often frequent a particular area for a period of several
days or weeks and then abruptly change their pattern and move to another
location (Hogan 1975, Shane 1977, Shane and Schmidly 1978, Würsig 1978,
Irvine et al. 1981). Whether they are following a particular prey species or
whether other factors account for these changes is not known.
Bottlenose dolphins often move with shrimp boats and other vessels that can
supply food (Norris and Prescott 1961, Leatherwood 1975, Wells et ul. 1980,
Gruber 1981). This movement is temporally adjusted to fit with the schedules
of these boats.
In summary, coastal bottlenose dolphins move with concentrations of food,
move into shallow safe areas, move with or against the tide and show some
regular (but usually not strong) diurnal movement patterns. The overriding
theme is variability. These are large-brained social mammals that can learn
much from their environment and retain some knowledge certainly for life. It
is likely that they know particular areas very well and that they remember when
and where the best chances for finding prey are likely to be.
BEHAVIOR AND ACTIVITY PATTERNS
Dolphin behavior patterns have been studied in captivity and in the wild,
and the literature contains many anecdotal accounts of behavior observed under
both conditions.
Captive Behavior
Captive bottlenose dolphin behavior has been reviewed by Tavolga (1966)

and Caldwell and Caldwell (1972a, 6). Classic work on captive T. truncatus
behavior done at Marineland of Florida (formerly Marine Studios) by McBride
and Hebb (1948), McBride and Kritzler (1951), Tavolga and Essapian (1957)
and Essapian (1963) described major characteristics of social structure, sexual
behavior, calving and play. As noted earlier, captive bottlenose dolphins form
dominance hierarchies with adult males dominant over all other tank-mates.
Sexual behavior is common and includes homosexuality and masturbation,
particularly in juvenile males. Mating occurs in late winter and spring and,
during these times, an adult male spends prolonged periods of time with a
receptive female. Male courtship includes rubbing, mouthing, nuzzling, “S-
curve” posturing, jaw-clapping and yelping, and females often respond to the
male or initiate similar behaviors. Gestation lasts 12 mo and pregnant females
spend most of the time alone or with one other adult female during the latter
half of their pregnancies. The “auntie” dolphin often continues to swim with
46 MARINE MAMMAL SCIENCE, VOL. 2, NO, 1, 1986
the mother after the calf s birth. Females in labor are surrounded by their tank-
mates who exhibit extreme excitement. Healthy newborns usually swim to the
surface unassisted for their first breaths, but mothers attempt to lift stillborn
infants to the surface. Calves are weaned after 18 mo but sometimes remain
closely associated with their mothers for longer. Play-with other dolphins,
with other animals and with objects in the tank-is an important component
of captive behavior, particularly for calves and juveniles (Caldwell and Caldwell
1972a).
Epimeletic or care-giving behavior is common in T. truncatus (Caldwell and
Caldwell 1966). It includes females, both in captivity and in the wild, carrying
their own dead young for days or weeks, as well as healthy individuals sup-
porting sick, injured or dead mature dolphins (sometimes of different species,
in captivity) at the surface.
A variety of behaviors has been associated with aggression (Brown and Norris
1956, Norris 1967). Biting, teeth-raking, jaw-clapping, fluke-slapping and
ramming are some. Aggressive behavior may be displayed in the context of
establishing or maintaining dominance hierarchies, protecting young or de-
fending food items. Tooth scratches on a dolphin’s skin may be evidence of
aggressive or sexual encounters. Caldwell and Caldwell (1977) noted that head-
to-head ramming is a precopulatory rather than aggressive behavior in bottle-
nose dolphins.
Activity patterns are probably significantly affected by the captive environ-
ment. McBride and Hebb (1948) recorded sleep periods following feeding times
and more time spent sleeping (always facing into the current in the tank) at
night than during the day. In a study of vocal activity, Powell (1966) noted a
peak in vocalizations just before sunrise, with fewest vocalizations occurring
from 2100 to 0300 hours.
A few authors have provided quantitative measures of captive behavior.
Saayman et al. (1973) found a peak in eight precopulatory behaviors at midday
and in the early afternoon in captive T. “aduncus. ” Courtship and copulatory
behavior were characterized and quantified by Puente and Dewsbury (1976),
who observed one pair of bottlenose dolphins. They found that specific behav-
iors like yelping and head butting were associated with copulation more often
than with general social behavior. Defran and Pryor (1980) reported on the
frequency of occurrence of 55 “behavioral events” in numerous captive ceta-
ceans including the bottlenose dolphin.
Behavior in the Wild
Several approaches have been taken to the study of free-ranging T. truncatus

activity patterns, but generally investigators focus on temporal, seasonal, eco-
logical, spatial and environmental effects on behavior. No set of behavioral
terms has been used consistently although the definitions provided by each
author indicate that the same broad activities are usually being analyzed. Feed-
ing, traveling, social interactions and idling are the major categories of behavior
recorded by most authors observing free-ranging animals. Most types of feeding
discussed by different authors are reviewed by Leatherwood (1975). Traveling

(or swimming) generally refers to dolphins involved in persistent, directional
movement. Social interactions are usually categorized as “mating,” “play,”
“rubbing,” etc., but most authors describe all of these categories as being
intricately intertwined and probably serving social as well as sexual functions
(Tayler and Saayman 1973, Norris 1974, Wells 1984). “Idling” and “resting”
usually refer to dolphins engaged in slow movements generally lacking com-
ponents of the other types of behavior described here. “Milling” involves fre-
quent changes in heading and may be associated with feeding, socializing or
play, if rapid, or with resting or idling, if leisurely. True resting behavior has
never been as clearly identified in T. truncatus as it has in some other cetaceans
such as the Hawaiian spinner dolphin, Stenella longirostris (Norris and Dohl
19806, Norris et al. 1982).
All reports of free-ranging bottlenose dolphin behavior indicate that the
animals are active at night as well as during the day (Tayler and Saayman
1972, Hogan 1975, Shane and Schmidly 1978, Irvine et al. 1981, Gruber
1981). However, indications of nocturnal activity are usually based on auditory
clues (breathing and splashing), anecdotes from fishermen or the absence of
diel variation in movements or respiratory patterns as indicated by radio-tag
signals. Direct observations are needed to define the actual level and type of
activity at night.
The earliest quantification of free-ranging bottlenose dolphin behavior was
made by Saayman et al. (1973) in South Africa. These authors found that
“feeding” peaked in the early morning and late afternoon (first reported by
Tayler and Saayman 1972), and that “mating” began after the morning feeding
period and continued until the afternoon feeding period began. “Leaping” was
associated with both mating and feeding while “rubbing” began when mating
began, but, unlike mating, continued throughout the evening feeding period.
Shane (1977) and Shane and Schmidly (1978) analyzed T. truncatus be-
havior in southern Texas. Major activities were tested (using Chi-square) for
possible effects of month, time of day, tidal stage, weather conditions and
location. Traveling occupied a significantly greater proportion of all observed
behaviors in January-April and June than during other months, and feeding
(six types of f ee d’ing were defined) occupied about twice as much time in
August-December and May as in other months (Fig. la). These feeding peaks
were associated with major migrations of fish and invertebrates in the fall and
spring. Socializing (= “mating”) peaked in April and May, corresponding with
a maximum of 10.6 percent calves in the population in the spring. Shane
(1977) found the same temporal pattern of feeding and mating (socializing)
(Fig. IS) reported by Saayman et al. (1973). She also noted that traveling
declined when resting (uniquely defined in this study as an animal maintaining
position against a strong current) increased in the evening. Tidal state signifi-
cantly influenced the frequency of both resting and bowriding (Fig. lc). Resting
occurred almost exclusively at ebb tide; this behavior could actually have been
feeding, with dolphins facing into the current and capturing fish carried out
with the tide (Shane 1977). Bowriding was generally restricted to ebb and flood
mate
feed
01 ’ ’ ’ ’ ’ ’ ’ ’ ’ ’ ’ ’
JJASONDJFMAM
MONTH
0700-0959 1000-1259 1300-1659 1700-1959

TIME PERIOD
-C
%-
bowride
4-
rest
0
ebb slack flood
TIDE
Figure 1. Relationship between bottlenose dolphin activity patterns and month, time
of day and tidal state near Aransas Pass, Texas. Only statistically significant relationships
(P < 0.005) are presented. The behavior labeled “mate” refers to all social behavior
including physical contact and frequent aerial behaviors.
tides; dolphins rode against the ebb tide 81 percent of the time (versus 19
percent of the time with it) and against the flood tide 57 percent of the time,
suggesting an energy conservation function for bowriding. Socializing (“mat-
ing”) and feeding were concentrated in particular areas during the study. So-
cializing (“mating”) took place in large open bays off the path of boat traffic
while feeding was most common at two 15-m deep “fishing holes.”
Wiirsig and Wiirsig (1979) discussed feeding, aerial and social behavior of
T. truncatus in Argentina. Dolphins appeared to rest near shore during the
morning. Aerial behavior was most frequent in the afternoon and included
leaps, headslaps, noseouts, tailslaps and kelp tossing. This behavior was often
accompanied by “nudging” between dolphins. Leaps were believed to be as-
sociated with feeding, whereas noseouts, belly-ups and kelp tossing were con-
sidered a part of play and copulatory behavior. The Würsigs hypothesized that
tailslaps, usually performed by one large adult, may have been warning signals
from a dominant animal. Widely spread groups of dolphins moving rapidly in
offshore waters were believed to be searching for food and “milling” upon the
discovery of a food source.
Gruber (1981) found an association between traveling, feeding and mating
and season and time of day for dolphins near Port O’Connor, Texas. Traveling
and mating occurred more frequently in spring and summer than in fall and
winter, but feeding occurred about 25 percent of the time in fall and winter
and only 5-10 percent of the time in spring and summer. Traveling and mating
increased gradually in frequency throughout the day, but feeding declined
throughout the day.
Most of the behavioral studies mentioned so far relied heavily on small boat
work. The researchers found that they could move slowly with groups for long
periods of time without appearing to greatly affect behavior. However, a cau-
tionary note was raised by Wiirsig and Wiirsig (1979, 1980), who monitored
bottlenose and dusky dolphins from cliffs and a small boat at the same time.
They found that, at times, a small vessel may herd the dolphins it is following
in a manner not perceived by observers on the boat. However, this effect was
not strong, and the dolphins moved in the same general direction as when the
boat was not present.
A few shorter-term studies provided additional or supportive information on
behavior. Hussenot (1980) described the behavior of T. truncatus in Brittany.
Leaping was associated with play and greeting behavior. Dolphins were believed
to be resting when facing against a strong current and maintaining position
there (as Shane 1977 described). Diving repeatedly in one location was believed
to indicate feeding. Tyack (1976) recorded acoustic emissions of Argentine
dolphins while describing their behavior. He was able to identify 15 categories
of sounds occurring during five activities and found significant differences in
types and frequencies of vocalization during the different activities. Hogan
(1975) reported on feeding, mating and play in bottlenose dolphins living near
the Georgia-South Carolina border. Mating and play were seen most often in
large aggregations in late summer and early fall and not seen at all in April.
No other seasonal patterns of behavior were noted.
50 MARINE MAMMAL SCIENCE, VOL. 2. NO. 1. 1986
Obviously lacking from the behavioral data on free-ranging bottlenose dol-

phins are observations made underwater. The tendency for bottlenose dolphins
to inhabit murky coastal waters often precludes underwater study.
Feeding
Bottlenose dolphin feeding behavior is nearly as diverse as the diet upon

which they feed (see Leatherwood 1975 for a summary of food types) (Caldwell
and Caldwell 1972a). Leatherwood ( 197 5) discussed three types of feeding in
association with shrimp boats, two types of cooperative feeding, “crowding” of
fish against shoals and shorelines and individual shallow-water feeding. The
shrimper-dolphin association is discussed in the Human-Dolphin Interactions
section ofthis paper. Hoese (1971), Hogan (1975) and Bel’kovich et al. (1978)
described dolphins trapping fish against the shoreline. Irvine et al. (1981)
observed rapid, individual shallow-water fishing and possible cooperative feed-
ing, as did Shane and Schmidly (1978). Tayler and Saayman (1972) also
described two cases of cooperative feeding by bottlenose dolphins. The following
case is thought to be representative of this pattern in many places. “Bottlenose
dolphins, leaping and splashing . . . converged slowly from opposing flanks and
gradually a tightly packed shoal of fish became discernible as a dark-coloured
mass beneath the surface between the encircling dolphins. Simultaneously, dol-
phins could be seen darting under the shoal and thus preventing it from sound-
ing. The shoal was consumed from the sides and underneath while the whole
ensemble progressed slowly out of sight at about 7 k.p.h.” (Tayler and Saayman
1972, p. 43). Bel’kovich et al. (1978) and Morozov (1970) gave detailed
descriptions of cooperative and individual feeding techniques used by T. trun-
catus in the Black Sea. As Wiirsig and Wiirsig (1980) found for dusky dol-
phins, (Lagenorhynchus o6sc~~z~s), they found certain types of leaps associated
with feeding behavior. A number of authors mention fish tossing: some ascribe
it to play (Gunter 1942, Shane and Schmidly 1978) while others suggest that
it may serve to soften or behead the fish (Norris and Prescott 1961, Wiirsig
and Wiirsig 1979) or simply to turn the fish so it can be ingested headfirst
(Bel’kovich et al. 1978). Leatherwood (1975), Bel’kovich et al. (1978) and
Barham et al. (1980) described dolphins pursuing fish while belly-up, possibly
to get the most direct line on the prey with their echolocation.
The bottlenose dolphin’s behavioral flexibility is most apparent in the diver-
sity of its feeding techniques. Dolphins take advantage of any readily available
food source, and they adapt their feeding methods according to food type and
local conditions. Some feeding methods become local traditions and are prob-
ably learned by succeeding generations (e.g., mud bank feeding in the Carolinas
or feeding behind shrimpers in the Gulf of Mexico).
Aerial and Boat Survey~
Very broad categories of behavior are frequently recorded for each sighting
of dolphins made during aerial or boat censuses. Observer differences and the
brief time spent on behavioral observations make these data of limited value,
but they are summarized in Table 2 for the sake of thoroughness.
Summa y
The behavioral data available on free-ranging bottlenose dolphins comes from

widely disparate locations and habitats. However, several broad generalizations
may be made: (1) dolphins appear to be active to some extent during both day
and night; (2) based upon diurnal observations, there are usually feeding peaks
in early morning and late afternoon; (3) the proportion of time devoted to
feeding seems to increase in the fall and winter, at least in Texas; (4) feeding
strategies are flexible and adapted to the habitat and food resources available;
and (5) social behavior is a major component of the animals’ daily activity
regime.
PREDATION
Sharks are probably the most important predator of bottlenose dolphins

worldwide, but killer whales are considered predators in some regions, such as
Argentine waters (Wiirsig and Wiirsig 1979). Even so, few data on shark
attacks on dolphins are available. From 3-18 percent of dolphins captured or
examined in three studies in Florida and South Africa bore definite shark-bite
scars (Wood et al. 1970, Ross 1977, Wells et al. 1980). However, shark-bite
scars are of uncertain value as indicators of levels of predation since they merely
represent unsuccessful predation attempts or the results of agonistic encounters
with sharks (e.g., sharks defending territory) at some unknown time during the
animals’ lives. Dolphin remains have been found in the stomachs of numerous
shark species, but most often in tiger (Galeocerdo cuvier), dusky (Carcharhinus
obscurus) and bull sharks (Carcharhinus leucas).
The responses of bottlenose dolphins to the presence of potential predators
vary relative to the species and size of the predator, the activity and size of the
dolphin group and the physical habitat. These responses include tolerance, active
avoidance and active aggression. Tiger sharks elicited a more marked reaction
by dolphins than did other shark species placed in a community tank (McBride
and Hebb 1948). In another study, a bottlenose dolphin conditioned to repel
various species of sharks responded appropriately to commands to repel sandbar
(Carcharhinus milberti), lemon (Negaprion brevirostris) and nurse (Ginglyostoma
cirratum) sharks, but when tests involved bull sharks the dolphin became
agitated and refused to respond to commands in repeated tests (Irvine et al.
1973).
Mutual tolerance between bottlenose dolphins and sharks has been reported
both from captivity (Essapian 1953, Wood et al. 1970) and the wild (AIBS
1967, Wood et al. 1970, Wells, pers. obs. 1975, Leatherwood, 1977). No
agonistic interactions were seen when pairs of bottlenose dolphins were put in
tanks with one to three individuals of various sizes of sandbar, lemon or bull
sharks (AIBS 1967, Gilbert et al. 1971, Irvine et al. 1973).
Table 2. Percentage of occurrence of different categories of behavior reported during four aerial survey studies and one boat study (Odell and
Asper 1982) of bottlenose dolphins.
Lear and Bryden

1980 Odell and Asper
1982
Leatherwood Barham et al. Rocky Ocean Odell and
1979 1980 nearshore beach Reynolds 1980 Unmarked Marked
Feeding (%) 36 49 8 18 15 21
Traveling (%) - 37 23 3; - 48 34
Idling/milling (%) - - 56 35 - 26 29
Mating/playing (%) - - 14 19 - 4 10
Sample size 64 groups 97 herds 2,698 individuals 326 herds 489 herds 269 herds
Time of survey Aug. 1977 Mar. 1978 1 year 1975-1976 Dec. 1980-Mar. 1982
Survev location E. Florida S. Texas E. Australia W. Florida E. Florida
SHANE El-AL.: BOTTLENOSE DOLPHINS: A REVIEW 53
Greater group cohesion and active avoidance have been reported as responses
to predators. Captive dolphins swam more rapidly and formed tighter groups
when sandbar sharks were introduced into their tank (McBride and Hebb 1948,
Wood et al. 1970). On another occasion all of the bottlenose dolphins in a
tank gathered around a female giving birth and herded approaching sharks
away (McBride and Hebb 1948). Free-ranging bottlenose dolphins off South
Africa actively avoided hammerhead (Sphyrna zygaena) and great white (Car-
charodon carcharias) sharks (Tayler and Saayman 1972).
A number of authors have reported aggression by dolphins towards sharks.
Bottlenose dolphins have attacked sharks in captivity (McBride and Hebb
1948, Essapian 1953, Brown and Norris 1956, Norris and Prescott 196 1) and
in the wild (Gunter 1954). Only one second-person account of a coordinated
agonistic response of bottlenose dolphins towards a shark in the wild has been
reported: Gunter (1942) received a report of dolphins killing a shark off Texas.
As discussed earlier, the relationship between group size and habitat char-
acteristics may be an important factor in dealing with predation pressure on the
west coast of Florida (Wells et al. 1980). Significantly smaller groups of dol-
phins were found in the complex, shallower inshore habitat than in the more
open Gulf of Mexico and Tampa Bay waters. It appears that, as the physical
habitat provides less protection in terms of reducing the volume of water that
must be monitored or providing physical barriers or paths for predators, the
importance of the group as a means of protection of dolphins from predators
may increase. More observations are necessary, however, to test this hypothesis.
HUMAN-DOLPHIN INTERACTIONS
Interactions between humans and dolphins range from apparently beneficial

to directly destructive to the animals.
Perhaps the most common association is between bottlenose dolphins and

boats. Dolphins are frequently attracted to the pressure waves created at the
bows of ships, and they bowride there (Norris and Prescott 1961, Walker
1975, Shane and Schmidly 1978, Gruber 1981). Bowriding may be a form
of play or it may be an energy-saving means of locomotion. Ship bowriding is
probably derived from whale bowriding. T. truncatus has been seen taking
advantage of the pressure waves at the heads of right whales (Würsig and
Würsig 1979), humpback whales (Darling, pers. comm.) and gray whales
(Leatherwood 1974). Dolphins also ride in the stern wake of vessels. A related
behavior is surfing, first reported by Caldwell and Fields (1959) and also seen
by Norris and Prescott (196 l), Saayman et al. ( 1973) and by the authors in
Texas, California, Hawaii and Argentina. Bottlenose dolphins frequently ride
waves alongside human surfers. On rare occasions a dolphin is killed by a ship’s
propeller (Shane and Schmidly 1978). Norris and Prescott (196 1) reported
that bottlenose dolphins regularly fed in San Diego Bay where Navy boats
dumped garbage. Norris (1974) reported bottlenose dolphins following ferries

in San Diego Bay, presumably to feed on organisms stirred up in the propeller
wash; Shane (1977) reported the same behavior in Texas. Hussenot (1980)
discussed different groups of dolphins in Brittany: some were attracted to boats
and accompanied them and others fled at the sound of motors. Wiirsig and
Würsig (1979) observed occasional tailslapping in response to the approach of
their boat and interpreted it as an indication of disturbance. Shane and Schmidly
(1978) and Wells (1978) reported that dolphins sometimes became attracted
to their research boats and often approached and accompanied them.
Capture
Bottlenose dolphins, being the most widely kept cetaceans in captivity, have
been captured by the hundreds in waters of the United States and other countries
(Odell et al. 1975, Walker 1975, Ridgway and Benirschke 1977, Leatherwood
and Reeves 1982, Perrin 1984). It is common knowledge that dolphins fre-
quently recognize boats previously used for capture and avoid them (Norris
and Prescott 1961, Irvine and Wells 1972, Leatherwood 1974, Norris 1974,
Norris and Dohl 1980a). However, Irvine et al. (1981) reported that dolphins
did not seem to avoid their “tagging-observation boat”; perhaps because it was
camouflaged and towed by another boat during tagging, the dolphins did not
associate the sound of its engine with the capture process.
Fisheries
Tursiops interacts with a number of fisheries, but none so much as the shrimp
fishery. Leatherwood ( 1975) delineated three major types of feeding in associ-
ation with shrimpers: (1) feeding behind actively trawling shrimp boats; (2)
feeding on “trash fish” discarded after a trawl; (3) feeding around non-working,
anchored shrimp boats. Dolphin/shrimp fishery interactions are also addressed
by Gunter (1942, 1954), Norris and Prescott (1961), Caldwell and Caldwell
(1972a), Hogan (1975), Shane and Schmidly (1978) and Wells etal. (19816).
Gruber (198 1) conducted interviews with Texas shrimp fishermen and observed
dolphin-shrimp boat interactions. Her observations concurred with those of
Gunter (1954) and Norris and Prescott (196 l), who noted a response to
changes in engine and winch sounds indicative of various stages of the shrimping
operation. Gruber (198 1) noted that dolphins moved toward shrimp boats at
least 1.5 km away, while Norris and Prescott (1961) saw dolphins approach
shrimpers from 2 mi away. Gruber found an increase in dolphin abundance in
one area at the time that shrimping began there, but she was unable to deter-
mine whether shrimping attracted more dolphins to the area or whether a
seasonal increase in the abundance of food there drew additional dolphins to
the area. Gruber’s interviews indicated that dolphins were infrequently drowned
in trawls (77 percent of the respondents had never captured a dolphin) and
that shrimpers usually had a benevolent attitude towards dolphins (although
23 percent had shot at dolphins and 18 percent had killed them). Some
shrimpers contended that dolphins tore their nets while taking fish from them
while others were certain that only sharks tore the nets and that dolphins
removed fish daintily (also recorded by Shane and Schmidly 1978). Irvine
(1975 in Odell et al. 1975) contended that fishermen on the west coast of
Florida generally “feel protective of local dolphins,” but evidence of net scarring
and a few dolphins killed by shooting showed that some conflicts occurred.
Busnel ( 197 3) described a “symbiotic’ ’ interaction between bottlenose and
humpbacked dolphins (Sousa teuszi) and native fishermen on the coast of
Mauritania fishing for mullet. The fishermen set up nets and beat the water to
“call” the dolphins; the dolphins herded fish against the nets, presumably
catching some for themselves while others were entrapped in the nets. In Geor-
gia (Dean 1979) and Florida (Shane, pers. obs.) crab fishermen fed dolphins
by hand from their boats.
Bottlenose dolphins have been reported to be a nuisance to at least three.
fisheries. In the Indian River in Florida, Orr (1976) and Leatherwood (1979)
reported damage by dolphins to lines and nets used in the mackerel fishery. To
our knowledge, these charges have never been investigated to the extent that
dolphins can be specifically implicated. In Florida, Cato and Prochaska (1976)
implicated T. truncatus in damaging handlines and nets. The theft of bait and
catch by bottlenose and rough-toothed dolphins (Steno bredanensis) from lines
in Hawaii has become a serious detriment to fishing there (Schlais 1984). Kuljis
et ul. (198 1) reported on tests of lithium chloride as aversive to dolphins when
inserted in bait; the substance failed to cause one-trial aversion but was consid-
ered to have potential for diverting dolphins from fishery predation. At Iki
Island, Japan, bottlenose dolphins and other small cetaceans are slaughtered by
fishermen who consider the animals detrimental to local fish stocks (Imanishi
1981). Incidental mortality occurs in fishing nets off west Florida (Wells et al.
1981a).
Contact
Direct association between humans and dolphins has occurred for a long
time and is described in tales told in ancient Greece (Devine and Clark 1967).
In recent times, bonds have been established between humans and bottlenose
dolphins under a variety of circumstances. Free-ranging individual dolphins
have been solicitous of interactions with humans in New Zealand (Alpers
1961), England (Dobbs 1977, Lockyer 1978, Webb 1978), South Africa
(Saayman and Tayler 1971, Tayler and Saayman 1972), France (Hussenot
1980) and the United States (Wells, unpubl.). In a number of cases, captive
dolphins have been released, but they have voluntarily remained at the release
site or subsequently initiated interactions with other humans (Cousteau and
Diole 1975). In one example, a male dolphin has remained in an enclosure at
a restaurant on Longboat Key, in Florida, for 15 yr. This animal remains in
spite of easy access to open water during all tides and the fact that free-ranging
dolphins frequently pass just outside of the enclosure (Wells and Shane, un-
publ.). In western Australia, over the past 20 yr a group of as many as seven
56 MARINEMAMMALSCIENCE,VOL. 2, NO. 1, 1986
bottlenose dolphins has regularly come into shallow water near shore to be fed
and petted (Gawain 1981, 1984, Connor and Smolker 1985).
Habitat Alteration
The impact of habitat destruction on bottlenose dolphins has not been thor-
oughly investigated. Odell (1976) suggested that pollution and heavy boat
traffic in Biscayne Bay may have contributed to an apparent decline in abun-
dance there. Reductions in bottlenose dolphin abundance in San Diego Bay
were linked to pollution (FAO-ACMRR 1978, p. 112), but Leatherwood and
Reeves (1982) contend that dolphins have returned to the bay since water
quality improved. Gunter (1942) believed that the bottlenose dolphin popu-
lation in south Texas declined since the early 1900s. Although no accurate
abundance estimates are available, popular opinion indicates that numbers have
declined in Galveston Bay concurrently with its increased use as a shipping
route and with increased pollution (Shane, unpubl.). Bottlenose dolphins are
extremely abundant in Aransas Pass despite its use by oil tankers and a large
number of smaller boats (Shane and Schmidly 1978). Extremely high chlori-
nated hydrocarbon residues have been found in the tissues of marine mammals,
including T. truncatus (O’Shea et al. 1980). In fact, the blubber of a California
specimen contained some of the highest PCB and DDE residues ever found in
cetaceans. DDT in California cetaceans and pinnipeds was the “highest known
for any populations of wild mammals” (O’Shea et al. 1980, p. 44). Habitat
alteration in the form of artificial passes in southern Texas may have opened .
up new habitat according to Leatherwood and Reeves (1983).
Human-dolphin interactions may have both positive and negative impacts
on dolphin populations. The bottlenose dolphin is certainly one of the most
adaptable of all marine mammals; Leatherwood and Reeves (1982) compare
it with the coyote and the white-tailed deer in this respect. Even so, habitat
alteration could potentially disrupt the social behavior, food supply and health
of these animals.
ACKNOWLEDGMENTS
The U.S. Fish and Wildlife Service and the University of Southern Mississippi provided
support for the preparation of a report upon which this paper is based. We thank Carol
King of the Moss Landing Marine Laboratories for typing the manuscript. G. Smith and
an anonymous reviewer provided helpful suggestions for the improvement of the paper.
LITERATURE CITED
AIBS, 1967. Conference on the shark-porpoise relationship, 9 November 1965. Amer-

ican Institute of Biological Science, Washington, DC.
ALPERS, A. 1961. Dolphins: the myth and the mammal. Houghton-Mifflin, Boston,
MA. 268 pp.
ASPER, E. D., AND D. K. ODELL. 1980. Bottlenose dolphin local herd monitoring:
capture, marking, collection of biological data, and follow-up observations of marked
animals. Final Report to National Marine Fisheries Service, Contract NA79-6A-C-

00027. Hubbs/Sea World Research Institute, 7007 Sea World Drive, Orlando,
FL 32809.
BARHAM, E. G., J. C. SWEENEY, S. LEATHERWOOD,R. K. BEGGS AND C. L. BARHAM.
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Received: April 11, 1985
Accepted: October 22, 1985

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MARINE MAMMAL SCIENCE, 2(1):34-63 (January 1986)

ECOLOGY, BEHAVIOR AND SOCIAL

The authors review the literature on bottlenose dolphin ecology, behavior

HOME RANGE AND STOCK DISCRETENESS

The concensus is that there is one species of Tursiops worldwide, separated

A home range is an area regularly used by an individual or group in the

reported that bottlenose dolphins found in the Molène archipelago of Brittany

It is only within the past lo-15 yr that information contributing to the

Table 1. Bottlenose dolphin density reported in various studies.

between populations of bottlenose dolphins in the Pacific Ocean on the basis

The patterns of social organization of bottlenose dolphins appear to be quite

Social Structure: Captive Animals

Since 1940 studies of social behavior at a number of oceanaria have produced

Social Structure: Free-Ranging Animals

While the nature of the social organization of bottlenose dolphins remains

SEASONALAND DAILY MOVEMENTS

While bottlenose dolphins may at times migrate in the higher-latitude ends

Diurnal and other short-term movement patterns appear to be influenced by

BEHAVIOR AND ACTIVITY PATTERNS

Captive bottlenose dolphin behavior has been reviewed by Tavolga (1966)

Behavior in the Wild

Several approaches have been taken to the study of free-ranging T. truncatus

discussed by different authors are reviewed by Leatherwood (1975). Traveling

0700-0959 1000-1259 1300-1659 1700-1959

Obviously lacking from the behavioral data on free-ranging bottlenose dol-

Bottlenose dolphin feeding behavior is nearly as diverse as the diet upon

Aerial and Boat Survey~

The behavioral data available on free-ranging bottlenose dolphins comes from

Sharks are probably the most important predator of bottlenose dolphins

Lear and Bryden

Interactions between humans and dolphins range from apparently beneficial

Perhaps the most common association is between bottlenose dolphins and

dumped garbage. Norris (1974) reported bottlenose dolphins following ferries

AIBS, 1967. Conference on the shark-porpoise relationship, 9 November 1965. Amer-

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